Colin M. Donihue

Adaptive evolution in urban ecosystems

Urban ecologists have demonstrated that cities are functioning ecosystems. It follows then that species living in these contexts should participate in and experience the same suite of biological processes, including evolution, that have occupied scientists for centuries in more “natural” contexts. In fact, urban ecosystems with myriad novel contexts, pressures, and species rosters provide unprecedentedly potent evolutionary stimuli. Here, we present the case for studying adaptive evolution in urban settings. We then review and synthesize techniques into a coherent approach for studying adaptive evolution in urban settings that combines observations of phenotypic divergence, measurements of fitness benefits of novel genetically based phenotypes, and experimental manipulations of potential drivers of adaptation. We believe that studying evolution in urban contexts can provide insights into fundamental evolutionary biology questions on rate, direction, and repeatability of evolution, and may inform species and ecosystem service conservation efforts.

Feed or fight: testing the impact of food availability and intraspecific aggression on the functional ecology of an island lizard

Summary 1. Body size often varies among insular populations relative to continental conspecifics – the ‘island rule’ – and functional, context-dependent morphological differences tend to track this body size variation on islands. 2. Two hypotheses are often proposed as potential drivers of insular population differences in morphology: one relating to diet and the other involving intraspecific competition and aggression. We directly tested whether differences in morphology and maximum bite capacity were explained by interisland changes in hardness of both available and consumed prey, and levels of lizard-to-lizard aggression among small-island populations. 3. Our study included 11 islands in the Greek Cyclades and made use of a gradient in island area spanning five orders of magnitude. We focused on the widespread lizard Podarcis erhardii. 4. We found that on smaller islands, P. erhardii body size was larger, head height was larger relative to body size, and maximum bite capacity became proportionally stronger. 5. This pattern in morphology and performance was not related to differences in diet, but was highly correlated with proxies of intraspecific aggression – bite scars and missing toes. 6. Our findings suggest that critical functional traits such as body size and bite force in P. erhardii follow the predictions of the island rule and are changing in response to changes in the competitive landscape across islands of different sizes.

Aegean wall lizards switch foraging modes, diet, and morphology in a human‐built environment

Abstract Foraging mode is a functional trait with cascading impacts on ecological communities. The foraging syndrome hypothesis posits a suite of concurrent traits that vary with foraging mode; however, comparative studies testing this hypothesis are typically interspecific. While foraging modes are often considered typological for a species when predicting foraging‐related traits or mode‐specific cascading impacts, intraspecific mode switching has been documented in some lizards. Mode‐switching lizards provide an opportunity to test foraging syndromes and explore how intraspecific variability in foraging mode might affect local ecological communities.Because lizard natural history is intimately tied to habitat use and structure, I tested for mode switching between populations of the Aegean wall lizard, Podarcis erhardii, inhabiting undisturbed habitat and human‐built rock walls on the Greek island of Naxos. I observed foraging behavior among 10 populations and tested lizard morphological and performance predictions at each site. Furthermore, I investigated the diet of lizards at each site relative to the available invertebrate community.I found that lizards living on rock walls were significantly more sedentary—sit and wait—than lizards at nonwall sites. I also found that head width increased in females and the ratio of hindlimbs to forelimbs in both sexes increased as predicted. Diet also changed, with nonwall lizards consuming a higher proportion of sedentary prey. Lizard bite force also varied significantly between sites; however, the pattern observed was opposite to that predicted, suggesting that bite force in these lizards may more closely relate to intraspecific competition than to diet.This study demonstrates microgeographic variability in lizard foraging mode as a result of human land use. In addition, these results demonstrate that foraging mode syndromes can shift intraspecifically with potential cascading effects on local ecological communities.

Microgeographic variation in locomotor traits among lizards in a human-built environment

Microgeographic variation in fitness-relevant traits may be more common than previously appreciated. The fitness of many vertebrates is directly related to their locomotor capacity, a whole-organism trait integrating behavior, morphology, and physiology. Because locomotion is inextricably related to context, I hypothesized that it might vary with habitat structure in a wide-ranging lizard, Podarcis erhardii, found in the Greek Cyclade Islands. I compared lizard populations living on human-built rock walls, a novel habitat with complex vertical structure, with nearby lizard populations that are naive to human-built infrastructure and live in flat, loose-substrate habitat. I tested for differences in morphology, behavior, and performance. Lizards from built sites were larger and had significantly (and relatively) longer forelimbs and hindlimbs. The differences in hindlimb morphology were especially pronounced for distal components—the foot and longest toe. These morphologies facilitated a significant behavioral shift in jumping propensity across a rocky experimental substrate. I found no difference in maximum velocity between these populations; however, females originating from wall sites potentially accelerated faster over the rocky experimental substrate. The variation between these closely neighboring populations suggests that the lizards inhabiting walls have experienced a suite of trait changes enabling them to take advantage of the novel habitat structure created by humans.

Hurricane-induced selection on the morphology of an island lizard

Hurricanes are catastrophically destructive. Beyond their toll on human life and livelihoods, hurricanes have tremendous and often long-lasting effects on ecological systems1,2. Despite many examples of mass mortality events following hurricanes3–5, hurricane-induced natural selection has not previously been demonstrated. Immediately after we finished a survey of Anolis scriptus—a common, small-bodied lizard found throughout the Turks and Caicos archipelago—our study populations were battered by Hurricanes Irma and Maria. Shortly thereafter, we revisited the populations to determine whether morphological traits related to clinging capacity had shifted in the intervening six weeks and found that populations of surviving lizards differed in body size, relative limb length and toepad size from those present before the storm. Our serendipitous study, which to our knowledge is the first to use an immediately before and after comparison6 to investigate selection caused by hurricanes, demonstrates that hurricanes can induce phenotypic change in a population and strongly implicates natural selection as the cause. In the decades ahead, as extreme climate events are predicted to become more intense and prevalent7,8, our understanding of evolutionary dynamics needs to incorporate the effects of these potentially severe selective episodes9–11.Two populations of Anolis lizards that survived Hurricanes Irma and Maria had larger toepads, longer forelimbs and shorter hindlimbs relative to the pre-hurricane populations, which suggests hurricane-induced natural selection.